The formation of the nestmate recognition template in ants and its effect on behavioural variation

我们对他人的行为取决于我们是否认识他们，例如通过他们的面孔。我们将感知到的面孔与存储在我们记忆中的原型面孔进行比较：模板。虽然识别的其他方面（生产组件，如提示表达;动作组件，如决策规则）得到了很好的研究，但感知组件，其中模板是一个基本方面，没有。目前，我们几乎不了解任何系统，人类或动物，存储在模板中的信息，以及模板如何形成。社会昆虫是一个特别好的模型来研究模板的形成，因为工人不同，他们对入侵者的侵略和我们表明，感知组件是负责的一些variation.Traditional观点的巢友识别模板描述它作为一个代表性的“平均群体气味”，这是相同的所有巢友。然而，模板可以是部分自指的，例如个体自身气味和平均群体气味的组合（"纽维模板"）。这可以解释观察到的行为差异：模板，因此识别，会随着每个人自己的气味而变化。对于模板的形成，也有两种替代假设：模板可以通过习惯化形成：蚂蚁“习惯”自己群体的气味。或者，联想学习也可以发挥作用：蚂蚁可能会将群体气味与与其他蚂蚁互动中的好或坏经验联系起来，如果个体经历不同的互动，这可能会导致个体间的差异。这将是特别明显的，如果蚂蚁学会关联非巢气味的负面经验（如打架），这将提高入侵者的检测从熟悉的非巢殖民地只有在那些个人，以前遇到了非巢殖民地的问题。我们从讨厌的邻居效应的实验中获得了后者的初步证据，在这个实验中，对非巢友的熟悉增加了蚂蚁对特定非巢友群体的攻击性。为了测试自我参照模板或模板的联想学习是否对观察到的巢友识别变化负责，我们将操纵蚂蚁工蚁的气味相对于它们的群体气味，并操纵个体所经历的相互作用。我们还将直接阻碍药物干预的联想学习。最后，我们将分析大脑和触角的基因表达，以跟踪我们的实验操作引起的分子效应。然后，我们将这些影响与不同的工人变体之间的分子差异进行比较，这些变体在它们的攻击性方面有所不同。我们的项目将揭示长期存在的，目前热议的问题，如何形成巢识别模板，与我们的理解行为变异和识别过程的一般影响。

Our behaviour towards others depends on whether we recognise them, for example by their faces. We compare the perceived face with prototypes of faces stored in our memory: the templates. While other aspects of recognition (production component, e.g. cue expression; action component, e.g. decision rules) are well studied, the perception component, of which template is a fundamental aspect, is not. Currently, we have little understanding from any system, human or animal, of the information stored in templates, and how templates are formed. Social insects are a particularly good model to study template formation because workers differ in their aggression towards intruders and we showed that the perception component is responsible for some of the variation.Traditional views of the nestmate recognition template describe it as a representation of the “average colony odour”, which is identical for all nestmates. However, the template could be partly self-referent, e.g. a combination of an individual's own odour and the average colony odour (“Newey template”). This would explain the behavioural variation that has been observed: Templates, and thus recognition, would vary with each individual's own odour. For template formation, there are two alternative hypotheses as well: The template could be formed by habituation: Ants “get used” to their own colony odour. Alternatively, associative learning could play a role: Ants may associate colony odours to good or bad experiences in interactions with other ants, which is likely to cause inter-individual variation if individuals experience different interactions. This would be particularly evident if ants learn to associate non-nestmate odours with negative experiences (e.g. fights), which would improve the detection of intruders from familiar non-nestmate colonies only in those individuals that previously encountered the non-nestmate colony in question. We have preliminary evidence for the latter from experiments on the nasty neighbour effect, where familiarity with non-nestmates increases an ant's aggression towards the specific non-nestmate colony.To test whether self-referent templates or associative learning of templates are responsible for the observed variation in nestmate recognition, we will manipulate the odours of ant workers relative to their colony odour and manipulate the interactions experienced by individuals. We will also directly impede associative learning with pharmacological interventions. Finally, we will analyse brain and antennal gene expression to track molecular effects caused by our experimental manipulations. We will then compare these effects to the molecular variation among different worker morphs, which vary in their aggression. Our project will shed light on the long-standing and currently hotly debated question of how nestmate recognition templates are formed, with implications for our understanding behavioural variation and recognition processes in general.